Geo log i cal Quar terly, 2017, 61 (1): 186–204 DOI: http://dx.doi.org/10.7306/gq.1312

Fa cies and fauna prox ies used to re con struct the MIS 5 and MIS 7 coastal en vi ron ments in east ern Tu ni sia

Amel CHAKROUN1, * and Dalila ZAGHBIB-TURKI1

1 Uni ver sity of Tu nis El Manar, De part ment of Ge ol ogy, Fac ulty of Sci ences of Tu nis, Uni ver sity Cam pus, 2092, El Manar II, Tu ni sia

Chakroun, A., Zaghbib-Turki, D., 2017. Fa cies and fauna prox ies used to re con struct the MIS 5 and MIS 7 coastal en vi ron - ments in east ern Tu ni sia. Geolog i cal Quarterly , 61 (1): 186–204, doi: 10.7306/gq.1312

This study is based on the anal y sis of the sedimentology and the fos sil as sem blages of Pleis to cene de pos its in East ern Tu ni - sia. It uses differ ent fauna assem blages to help infer palaeoenvironments and in par tic u lar dis cusses the im por tance of the dis tri bu tion of Persististrombus latus, es pe cially as re gards MIS 5e de pos its around the Med i ter ra nean Ba sin. The strati - graphic suc ces sion has been di vided into two main units. The lower unit is quartz-rich. Its basal part contains a marine fauna that includes well-preserved bi valves, gastro pods , echinoids and cirripedes domi nated by Acanthocardia, Cerastoderma, Parvicardium, Arca, Donax, Glycymeris, Loripes, Tellina and Cerithium. This fauna rep re sents a typ i cal as so ci a- tion/biocenosis typ i cally de vel oped in a shel tered bay, of mod er ate to low wave/cur rent ac tion and rapid post-mor tem burial. The up per unit is car bon ate-dom i nated. Its basal part also con tains a rich ma rine fauna that in cludes the ex tant trop i cal West Af ri can spe cies Persististrombus latus. The oc cur rence of Persististrombus latus in this unit com pared to its pres ence in the whole Med i ter ra nean ba sin in the Late Pleis to cene leads us to ques tion its im por tance in the in ter pre ta tion of cli mate warm - ing dur ing the Late Pleis to cene, and to sug gest its At lan tic ori gin .

Key words: MIS 5 and MIS 7, Tu ni sia, malacofauna, palaeoec ol ogy , Persististrombus latus, palaeoclimate.

INTRODUCTION ties help to re con struct their depositional en vi ron ments or biotopes de vel oped along the east ern coast of Tu ni sia.

Pleis to cene de pos its of the east ern coast of Tu ni sia cor re la - tive to ma rine iso tope stage (MIS) 5, MIS 7 and MIS 9 have been MATERIAL AND METHODS the sub ject of sev eral stud ies (Allemand-Mar tin, 1923, 1924; Castany, 1954, 1955, 1956; Barrot, 1972; Paskoff and Along the east ern Tu ni sian coast, from the Cap Bon to Sanlaville, 1976, 1977, 1978, 1980, 1983; Oueslati, 1980; Hergla ar eas, the Pleis to cene (MIS 5e and MIS 7) suc ces sion, Oueslati et al., 1982; Blanc, 1982; Bernat et al., 1985; Miller et out crop ping at the cit ies of Menzel Temime, Qorba and Hergla al., 1986; Mahmoudi, 1986, 1988; Oueslati, 1994; Jedoui, 2000; (Fig. 1), is in ves ti gated. Five sec tions across this suc ces sion Jedoui et al., 2001, 2002, 2003; Bouaziz et al., 2003; Le Guern, are de tailed (Fig. 2). In the study, the stra tig ra phy was doc u - 2005; Chakroun et al., 2005, 2009a, b, 2016; Chakroun, 2006; mented, and sam ples were col lected from the car bon ate and Mauz et al., 2009; Mejri et al., 2012). This suc ces sion, con tain ing the quartz-rich units (MT, Q2 and Hg1 sec tions) com pos ing this a ma rine fauna, has been doc u mented as transgressive. In east - suc ces sion. ern Tu ni sia, it is un der lain by a ma jor un con formity. It can di rectly We col lected a range of spe cies pres ent within the dif fer ent over lie con ti nen tal as well as ma rine Mio cene de pos its or the units. Ad di tion ally, we pro cessed (1) by ex trac tion of large spec - ma rine Plio cene, or even con ti nen tal Villafranchian de pos its. i mens of all taxa (larger than 2 mm in size) tak ing into ac count In the pres ent work we de tail the fos sil con tent of the MIS 5e their stratinomy and po si tion within the sed i ment, (2) by ex tract - and MIS 7 de pos its out crop ping along the Cap Bon Pen in sula ing 500 g of dry de pos its and gath er ing small spec i mens from and far ther to wards the south in the Hergla area. The fau nal spe - their res i dues ob tained in sieves (us ing a 1 mm mesh screen). cies dis tri bu tion and taphonomy are an a lysed. The main eco log i - All spec i mens that re mained on the screen were picked and cal char ac ter is tics of the dif fer ent in ver te brate spe cies com mu ni - iden ti fied and counted (Ta bles 1 and 2). Typ i cal spe cies as - sem blages are also de fined and any change in their as sem- blages com po si tion is de tected and com mented on. For tax o nomic iden ti fi ca tion and eco log i cal as sig na tion, we *Corresponding author, e-mail: [email protected] re ferred to Abbott (1960), Perrier (1954), Fekih (1975), Meco (1977) and Sessa et al. (2013). Tax o nomic at tri bu tions were Received: March 30, 2016; accepted: June 13, 2016; first published up dated by us ing the World Reg is ter of Ma rine Spe cies Da ta - online: September 12, 2016 Facies and fauna proxies used to reconstruct the MIS 5 and MIS 7 coastal environments in eastern Tunisia 187

green clay lay ers ob served only at Menzel Temime and (3) coarse sandy de pos its with dune cross-bed ding or densely packed root struc tures. Thus, to wards the top of this lower part a con ti nen tal char ac ter is de vel oped. The car bon ate unit is sig nif i cantly better de vel oped in the Cap Bon area as well as along the east ern coast of Tu ni sia. It con sists of 12 m of car bon ate de pos its, out crop ping at var i ous al ti tudes rang ing from 5 to 10 m above the pres ent sea level. It is com posed of two parts: 1. Centi metre- to metre-scale chalky sands rich in al tered ma rine mol lusc shells. These de pos its are or ga nized into four nor mally graded bed ding se quences un der lain by an ero sive sur face. This basal part re cords the ac tion of lo cal ep i sodic storms along this northeast ern coastal area. In this part are pre - served the main spe cies char ac ter iz ing the Sen e ga lese fauna, Persististrombus latus, a proxy of warm trop i cal ma rine wa ter. 2. Decimetric-scale car bon ate de pos its rich in ooliths and pel lets with dune cross-bed ding. This part con tains a con ti nen - tal in ver te brate fauna com posed mainly of helicid gas tro pods in clud ing He lix spp. and Rumina decollata. Lo cally, close to Qorba city, these helicids are as so ci ated with large mam mal spe cies: Phacochoerus aethiopicus, Gazella atlantica, Alcelaphus buselaphus, Equus mauritanicus, Ceratotherium simum and Hystrix cristata (Chakroun et al., 2005). To wards the south of Cap Bon, in the Sahel re gion close to the vil lage of Hergla (Fig. 1), there is an Up per Pleis to cene suc- ces sion that was stud ied by Mahmoudi (1988). This au thor dis - tin guished two for ma tions: the Kniss For ma tion and the Rejiche For ma tion, and he at trib uted this suc ces sion to the Tyrrhenian on the base of the oc cur rence of Persististombus latus (a se nior syn onym of Strombus bubonius). The basal part of the quartz-rich unit in the Hergla area (Hg1 and Hg2 sec tions) is equiv a lent to the Khnis For ma tion (Fig. 2). It is com posed from base to top of (1) fine sands, with rare crus- ta cean pel lets and ooliths, the fauna be ing rep re sented ei ther by de bris and small en tire bi valve shells, es pe cially cardiids and lucinids; (2) green clay and silts and (3) sandy de pos its with densely packed root struc tures. Fig. 1. Location map of the Tyrrhenian sites studied: The car bon ate unit that is equiv a lent to the Rejiche For ma- Menzel Temime, Qorba and Hergla tion is com posed of two parts: the lower part con sists of less or more con sol i dated car bon ate sands rich in molluscs and in clud - ing Persististrombus latus. The up per part is made up of sands base (http://www.marinespecies.org) and the Paleobiology Da- rich in helicids and root struc tures. ta base (https://paleobiodb.org/#/). CHRONOLOGICAL BACKGROUND FACIES AND GEOLOGICAL SETTING The best de vel oped Pleis to cene de pos its out crop ping in Along the east ern coast of Tu ni sia the Pleis to cene suc ces- east ern Tu ni sia have been at trib uted to the Tyrrhenian and sion ana lysed (MIS 5e and MIS 7) are far from ho mo ge neous. were char ac ter ized by the oc cur rence of Persististombus latus In our pre vi ous stud ies based on the de tailed anal y sis of stra tig- (Paskoff and Sanlaville, 1978). ra phy, lithofacies, morphoscopy, exoscopy, and palaeontolo - Ra dio met ric age de ter mi na tion has been made on the gical con tent (ver te brates and in ver te brates), we rec og nized Persististrombus latus-bear ing de pos its at dif fer ent lo cal i ties in two main Mid dle to Late Pleis to cene units, each one de fin ing a east ern Tu ni sia. U-se ries dat ing yielded re li able ages (Ta ble 3). com plete transgressive cy cle (Chakroun et al., 2005, 2009a, b; These ages (126 ± 7 Ky: Miller et al., 1986; 120 Ky: Jedoui et Chakroun, 2006). These two units are dis tin guished by their al., 2003) at trib ute the ma rine car bon ate mem ber of the Rejiche sedimentological and their palaeontological char ac ter is tics For ma tion to MIS 5e. This ma rine mem ber is over lain by de pos - (Fig. 2). Be low, a con cise de scrip tion of these units in the its with eolian cross-strat i fi ca tion at trib uted to MIS 4 (Mauz et Menzel Temime and Qorba sec tions is given. al., 2009). The lower unit, rarely de vel oped, is char ac ter ized by At Qorba and Hergla the out crops (Fig. 2) host the ma rine quartz-rich de pos its. It is com posed, from base to top, of (1) thin warm-wa ter gas tro pod Persististrombus latus in the car bon ate lay ers of fine sands rich in en tire shells of ma rine molluscs such unit, at trib ut ing it to the Rejiche For ma tion (MIS 5e). The ab- as Glycymeris nummaria, Loripes lucinalis, Ceratoderma edule, sence of Persististrombus latus in the car bon ate ma rine level of Parvicardium exiguum, Acanthocardia tuberculatum, Donax Menzel Temime make us less cer tain about the age of this lat ter trunculus, Tellina planata and Cerithium vulgatum; (2) len tic u lar de posit. How ever, the pres ence of the amphi-At lan tic spe cies 188 Amel Chakroun and Dalila Zaghbib-Turki s r e b m e m

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Ta ble 1

Taxonomic com posi tion and fre quency of spec i mens iden ti fied in the Mid-Up per Pleis tocene quartz-rich unit of Tu ni sia

MENZEL TEMIME QORBA HERGLA KHNIS (a) QUARTZ-RICH UNIT SPECIES NI [%] NI [%] NI [%] Acanthocardia tuberculata Linne 17 7 22 20 24 24 * Cerastoderma edule Linne 18 8 25 23 20 20 * Parvicardium exiguum Gmelin 22 10 23 21 20 20 Barbatia barbata Linne * Arca noae Linne * 17 16 2 2 *** Arca tetragona Poli 2 2 *** Pseudochama gryphina Lamarck * 2 Dosinia exoleta Linne * * Dosinia lupinus Linne 2 1 Donax trunculus Linne 31 14 Eastonia rugosa Helbling * * Gastrana fragilis Linne * *** Glycymeris nummaria Linne 12 5 17 16 2 2 Loripes lucinalis Lamarck 28 12 18 18 * Mactra stultorum Linne * * Mactra glauca Born * * Callista chione Linne * Pitar rudis Poli 2 1 Modiolus barbatus Linne * * Solecurtus strigilatus Linne * Pecten jacobaeus Linne 4 2 Ensis ensis Linne 5 2 Solecurtus strigilatus Linne 5 2 * Ruditapes decussatus Linne 2 2 Polititapes rhomboides Pen nant 4 2 2 2 Tellina planata Linne 33 14 * * Venus verrucosa Linne 5 2 3 3 Chamelea galina Linne 4 2 3 3 Bolma rugosa Linne * Neverita josephinia Risso * Hexaplex trunculus Linne * Semicassis granulata undulata Gmelin * 4 4 Cerithium vulgatum Bruguiere 19 8 * * Nassarius sufflatus Gould 5 2 * Nassarius reticulatus Linne 6 3 Cyclope neritea Linne 6 3 Balanus sp. * Echinocardium cordatum Pen nant * Bulk sam ple rich ness = 38 taxa 228 100 108 100 100 100

(a) – addi tional data from (Mahmoudi, 1986); * – spe cies identi fied from field survey of the quartz-rich unit: * – com mon; ** – very common; *** – abun dant; NI – number of in di vid uals (for bi valve taxa, this is the valve with higher frequency)

Stramonita haemastoma, which is gen er ally as so ci ated with This lat ter for ma tion has long been thought to date to MIS 5 Persististrombus latus at this level and its strati graphic po si tion (Tyrrhenian). Based on U/Th dat ing, Miller et al. (1986) corre - above the quartz-rich Douira For ma tion (MIS 7), sug gests that lated the marine quartz-rich Douira For ma tion with MIS 7. New the de pos its can be cor re lated with Rejiche For ma tion (MIS 5e). IRSL anal ysis of the Douira For ma tion, in the Cap Bon Pen in - Fur ther dat ing may yield a more pre cise age of this car bon ate sula, sup port the manty pre vi ous works (Miller et al., 1986; unit at Menzel Temime. Mauz et al., 2009; Mejri et al., 2012; Balescu et al., 2015). The ma rine quartz-rich deposit lack ing Persististrombus At the Khnis lo cal ity, the Khnis unit has an age of 121 latus, richly fossiliferous, is at trib uted to the Douira For ma tion. ± 10 Ky (Mauz et al., 2009). 190 Amel Chakroun and Dalila Zaghbib-Turki

Ta ble 2

Tax o nomic com po si tion and fre quency of spec i mens iden ti fied in the Mid-Up per Pleis to cene car bon ate unit of Tu ni sia

MENZEL TEMIME QORBA HERGLA KHNIS (a) CARBONATE UNIT SPECIES NI [%] NI [%] NI [%] Acanthocardia tuberculata Linne 2 3 22 19 36 20 * Cerastoderma edule Linne 3 4 34 19 Parvicardium exiguum Gmelin 2 3 25 14 Glycymeris nummaria Linne 12 17 24 24 29 16 * Arca noae Linne 4 4 29 16 * Arca tetragona Poli * * Pecten jacobaeus Linne 10 14 4 5 Spondylus gaederopus Linne * * * Ostrea edulis Linne 9 12 1 1 Loripes Lucinalis Lamarck 1 1 6 3 * Pseudochama gryphina Lamarck * Chama gryphoides Linne * * * Callista chione Linne * * Dosinia exoleta Linne * Dosinia lupinus Linne * Ruditapes decussatus Linne * Polititapes rhomboides Pen nant * Chamelea gallina Linne * Ve nus verrucosa Linne 1 1 * * Donax trunculus Linne 1 1 Eastonia rugosa Helbling 1 1 Mactra glauca Born * 6 3 Mactra stultorum Linne * * Callista chione Linne * * Gastrana fragilis Linne 1 1 Tellina planata Linne 1 1 6 3 * Diodora italica Defrance 2 2 * Diodora graeca Linne 1 1 Patella ferruginea Gmelin 1 1 Pa tella vulgata Linne * Patella caerula Linne 1 1 Naticarius stercusmuscarum Gmelin * Rissoa membranacea Ad ams * Tarantinaea lignaria Linne * Turritella communis Risso 3 3 * * Hexaplex trunculus Linne 14 20 2 2 * * Stramonita haemastoma Linne 12 17 2 2 * Cerithium vulgatum Bruguiere 10 10 Bittium reticulatum Da Costa 3 3 Monophorus perversus Linne 1 1 Cerithiopsis min ima Brusina * Cerithiopsis tubercularis Montag * Persististrombus latus Gmelin * * * Semicassis granulata undulata Gmelin 7 10 10 10 * * Nassarius sufflatus Gould * * * Nassarius reticulatus Linne 10 10 Cyclope neritea Linne * Columbella rustica Linne 4 4 * * Co nus ventricosus Gmelin 6 * Neverita josephinia Born * * Vermetus triquetrus Bivona Bernardi * * Thylacodes arenarius Linne * Bulk sam ple rich ness = 52 taxa 71 100 100 181 100

Ex pla na tions as in Table 1 Facies and fauna proxies used to reconstruct the MIS 5 and MIS 7 coastal environments in eastern Tunisia 191

Ta ble 3

Dat ing re sult sum mary of the Pleis to cene lit to ral de pos its of the Tu ni sian coast

For ma tions (Paskoff and Mahmoudi Pres ent Sanlaville, (1988) work Terms Lo cal ity Dat ing tech nique Age (Kys) MIS Ref er ences 1983) Ae olian OSL 68–122 4 Mauz et al. (2009) Rejiche Rejiche Car bon ate Monastir Th/U 126 ± 7 5e Miller et al. (1986) unit unit Ma rine Djerba Is land Th/U 120 5e Jedoui et al. (2003) Quartz-rich Khnis unit unit La goonal Khnis OSL 121 ± 10 5e Mauz et al. (2009) Menzel Temime Amino-acid data – 7 Miller et al. (1986) Douira OSL 138 ± 9 7 Mauz et al. (2009) Ma rine El Hajeb IRSL 216 7 Mejri et al. (2012) Douira Douira unit El Hajeb IRSL 333 9 Mejri et al. (2012) Hergla IRSL – ? Mejri et al. (2012) La goonal IRSL 182 ± 21 7 Balescu et al. (2015) Cap Bon Cap Bon IRSL 205 ± 16 7 Balescu et al. (2015)

RESULTS zoans or per fo rated by cliones or li chens; this sug gests that they were quickly bur ied where they lived. The fauna studied from the Pleis to cene successsion is very Be sides, the fau nal as sem blage men tioned above, from the di verse, dif fer ing from one unit to an other and be ing dis tin - quartz-rich unit of the MT sec tion con sist ing mainly of guished by as sem blage spe cies com po si tion and fre quency. endobenthic spe cies, was capa ble of be ing bur ied in soft sand The Up per Pleis to cene fos sils col lected in the dif fer ent units in or mud. There fore, such an as sem blage sug gests a shal - east ern Tu ni sia al low the de ter mi na tion of dif fer ent fau nal as - low-ma rine en vi ron ment, though quiet and lit tle af fected by cur- sem blages and the coastal sed i men tary en vi ron ments that they rents (e.g., tides, waves, swells, storms), far from wave ac tion, re flect. e.g., in shel tered bay. The pre ponder ance of molluscs con fined to soft sub strate is con sis tent with the grain size dis tri bu tion within the basal part of the quartz-rich unit out crop ping in the ASSEMBLAGES FROM THE MARINE PART OF THE QUARTZ-RICH UNIT Cap Bon Pen in sula (Chakroun, 2006). Among the stenohaline spe cies, echinoids are rare, though MENZEL TEMIME SECTION (MT) rep re sented by many dis so ci ated el e ments, e.g., spines and plates such as those at trib uted to Echinocardium cordatum, an In the thin shelly layer, the sandy basal part of the endobenthic spe cies. Today this taxon lives bur ied in sandy or quartz-rich unit (Fig. 3A), has been at trib uted to MIS 7 (Miller et silty sub strates, like most of the ma rine fauna rec og nized in the al., 1986). The ma rine mol lusc fauna iden ti fied here (Fig. 4) is ma rine part of the quartz-rich unit of the Menzel Temime sec- abun dant and in cludes 34 spe cies. It con tains mainly tion. Glob ally this endobenthic fauna may be consid ered as Acanthocardia tuberculata, Cerastoderma edule, Parvicardium autochthonous/subautochthonous and rep re sen ta tive of biota exiguum, Arca noae, Donax trunculus, Loripes lucinalis, Tellina char ac ter iz ing shal low but quiet ma rine-wa ter. planata and Cerithium vulgatum. Less abun dant spe cies are Dosinia lupinus, Glycymeris nummaria, Pitar rudis, Solecurtus QORBA SECTIONS (Q1 AND Q2) strigilatus, Pecten jacobaeus, Ensis ensis, Polititapes rhomboides, Ve nus verrucosa, Chamelea gallina, Nassarius In the Qorba re gion, be hind the pres ent-day shore line, near mutabilis, N. reticulatus, Cyclope neritea and Balanus sp. (Ta - the cave of El Geffel (Fig. 2, Q1 and Q2 sec tions), to the west, ble 1). the MIS 7 sand de pos its from the base of the suc ces sion stud - The shells of bi valves, es pecially those of Loripes lucinalis, ied is rich in fos sils rep re sen ta tive of a bi otic com mu nity domi - Tellina planata and Donax trunculus are of ten well-pre served, nated by Acanthocardia tuberculata, Parvicardium exiguum, in life po si tion, with their two contig uous valves. In this as sem - Cerastoderma edule, Glycymeris nummaria and Arca noae blage, the dom i nant ma rine spe cies are euryhaline. The (Ta ble 1). In deed, in this basal part of the quartz-rich unit the stenohaline spe cies such as those be long ing to Neverita and fauna is abun dant but less di verse (7 spe cies) and smaller than Hexaplex are rare. More over, in these shelly sands the spe cies in the Menzel Temime sec tion. Besides, the entire shells and di ver sity is high. This in di cates sta ble and fa vour able en vi ron - scarce de bris are not sorted but form reg ular subhorizontal thin men tal con di tions. layers with out any graded bedding (Fig. 3B). The well-pre - The dom i nance of en tire and well-pre served shells of many bi - served shells sug gest lit tle lat eral trans por ta tion of the fauna. valves be long ing to di verse gen era (e.g., Loripes, Acanthocardia, The as sem blage of this basal part of the quartz-rich unit con - Polititapes, Ve nus, Donax, Tellina) dem on strates that they have tains abun dant euryhaline molluscs (e.g., Cerastoderma not un der gone long trans por ta tion. Thus, they are in du bi ta bly fos - edule). The stenohaline spe cies, e;g., Semicassis granulata sil ised in or close to their appro pri ate biotope. Fur ther more, the undulata are rep re sented by few spec i mens (4%). In ad di tion shells of these molluscs are never en crusted by algae or bryo- most of them are endobenthic, liv ing bur ied in soft sub strate. 192 Amel Chakroun and Dalila Zaghbib-Turki

Fig. 3A – general view of the MT section; B – fossiliferous plane laminations of the quartzose marine member (Qorba); C – conglomeratic deposit of the carbonate marine member (Qorba); D – general view of Hg1 section Facies and fauna proxies used to reconstruct the MIS 5 and MIS 7 coastal environments in eastern Tunisia 193

Fig. 4Aa, b – Acanthocardia tuberculata, Ba, b – Parvicardium exiguum, Ca, b – Cerastoderma edule, Da, b – Glycymeris nummaria, E – Parvicardium exiguum, Fa, b – Arca noae, Ga, b – Donax trunculus, Ha, b – Dosinia exoleta, Ia, b – Venus verrucosa, Ja, b – Cerithium vulgatum, Ka, b – Nassarius sufflatus, La, b – Nassarius reticulatus 194 Amel Chakroun and Dalila Zaghbib-Turki

HERGLA SECTIONS (HG1 AND HG2 SECTIONS) the shal low Med i ter ra nean sea and along the west ern coast of Af rica from Mo rocco to Sen egal (Clench, 1947; Weisrock et al., In the Hg1 and Hg2 sec tions the basal part of the quartz-rich 1999), in shal low seas around the Ca nary Islands (Ramirez et unit (Fig. 2) is com posed of fine sands, with rare crus ta cean al., 2009) and the Azores (Spence et al., 1990). In the Med i ter - pel lets and ooliths, the fau nal as sem blage be ing rep re sented ra nean this spe cies seems to in habit sim i lar hab i tats to that of ei ther by debris or by small bi valve shells. It is dom i nated by Persististrombus latus (Rilov et al., 2001). The abundance of Acanthocardia tuberculata, Ceratoderma edule, Parvicardium Stramonita haemastoma on trop i cal coasts of Af rica and Amer - exiguum and Loripes lucinalis (Ta ble 1). Other spe cies are rare, ica shows that this spe cies re quires warm coastal ma rine wa - and in clude Arca noae, A. tetragona, Pseudochama gryphina, ters (But ler, 1985; Brown and Rich ard son, 1987). Near the Glycymeris nummaria, Ruditapes decussatus, Polititapes coast of Sen egal it is as so ci ated with strombids and defines the rhomboides, Ve nus verrucosa and Chamelea gallina. Sen ega lese fauna liv ing in shal low warm ma rine waters. Be - This fa cies of this basal part is over lain by a sandy level with sides, the abun dance of Stramonita haemastoma on trop i cal pla nar lami nations then by lentic ular clay with rare and sub trop i cal coasts of Af rica and Amer ica ad di tion ally in di - Cerastoderma edule. cates that this spe cies re quires warm shal low-ma rine waters (But ler, 1985; Brown and Rich ard son, 1987). ASSEMBLAGES FROM MIS 6 OR/AND MIS 4 Sec ondly, the pres er va tion state of this fauna (disarticulated AEOLIAN QUARTZ-RICH UNIT valves, shell de bris, worn shells) re flects strong coastal dy nam - ics, with wave and swell ac tiv ity. This is cor rob orated by the The depos its of the up per part of the quartz-rich unit, out - pres ence in this ma rine part of the car bon ate unit of angu lar crop ping spo rad i cally along the east ern coast of Tuni sia, are blocks of decimetre size and rounded peb bles. not very fossiliferous. In the Menzel Temime sec tions, they con- sist of sands which are very bioturbated by roots. In the Qorba QORBA SECTIONS sec tions this de posit is 1 to 2 m thick and it con tains both ma rine and con ti nen tal fos sils (e.g., He lix spp., Rumina decollata). This Tow ards the west, in the Q2 sec tion (Fig. 2) be hind the El as sem blage in di cates a la goon linked to the sea. Geffel cave, the fauna of the ma rine part of the car bonate unit is In the Hergla area, the con ti nental quartz-rich depos its (with both very abundant, very diverse (46 spe cies) and marked by an av er age thick ness of 1 m) are com posed of fine sands and the oc cur rence of Persististrombus latus, Semicassis gran u- pre serve cross-bed ding com monly destroyed by intense late, Stramonita haemastoma, Co nus ventricosus (Fig. 5). The bioturbation. They contain only a very few helicid shells. Locally as sem blage is dom i nated by Acanthocardia tuberculata and the top of the quartz-rich unit is marked by intense root struc - Glycymeris nummaria. tures and may be other wise unfossiliferous. The as so ci ated spe cies are nu mer ous (Ta ble 2): Arca noae, Pecten jacobaeus, Spondylus gaederopus, Ostrea ASSEMBLAGES FROM THE MARINE PART edulis, Loripes lucinalis, Pseudochama gryphina, Chama OF THE CARBONATE UNIT gryphoides, Callista chione, Dosinia exoleta, Do. lupinus, Ruditapes decussatus, Polititapes rhomboides, Chamelea The fau nal as sem blages in the lower term of the car bonate galina, Ve nus verrucosa, Donax trunculus, Eastonia rugusa, unit are dif fer ent from those of the un der lying unit. Mactra glauca, M. stultorum, Callista chione, Gastrana fragilis, Tellina planata, Diodora italica, D. graeca, Pa tella ferruginea, P. MENZEL TEMIME SECTION vulgata, P. caerula, Naticarius stercusmuscarum, Rissoa membranacea, Tarantinaea lignaria, Turritella communis, The basal term of the car bonate unit out crop ping in the Hexaplex trunculus, Stramonita haemasto ma, Cerithium Menzel Temime area (Fig. 2; MT sec tion, Fig. 3A) is rich in ma - vulgatum, Bittium reticulatum, Monophorus perversus, rine fos sils. The fauna is dom i nated by Glycymeris nummaria, Cerithiopsis minima , C. tubercularis, Persististrombus latus, Hexaplex trunculus, Stramonita haemastoma, Semicassis Semicassis granulata undulata, Nassarius mutabilis, N. granulata undulata, Pecten jacobaeus and Ostrea edulis (Ta - reticulatus, Cyclope neritea, Columbella rustica and ble 2). The bi valve valves are of ten stacked one above the Thylacodes arenarius. other to form a coquina with an av er age thick ness of 50 cm. Among gas tro pods, spe cies Cerithium vulgatum, Semicas- The shells are of ten poorly pre served (worn, cor roded, rolled sis granulata undulata and Nassarius reticulatus are the most and bro ken). In these de pos its, the <2 mm sized frac tion con - com mon. sists mainly of very fine debris of these shells, ooids and pel lets. This very diverse as sem blage (46 spe cies) likely reflect s a Iso lated quartz grains are rare; these grains as well as shell de - ma rine palaeoenvironment cor re spond ing to the photosyn thetic ++ - bris form the ooid nuclei. Ben thic foraminifera are rare; they are zone, in which Ca and CO3 ions were suf fi ciently avail able to com posed mainly by Am mo nia beccarii and Elphidium crispum. al low for ma tion of thick cal car e ous shells. The com po nents of The abun dant ooliths and the poor pres er va tion of shells re flect this biocenosis would be de pend ent on suf fi cient pri mary pro- more in tense ma rine dy nam ics than those char ac ter iz ing the duc ers and hence suf fi cient nu tri ents. ma rine part of the quartz-rich unit. All the char ac ter is tics of the As re gards the substrat e, the bi valves of the Veneridae fam - as sem blage from the basal term of the car bon ate unit suggest ily, e.g., Ve nus and Donax in di cate a soft sub strate com posed that the biota was not fos sil ised in situ but un der went sub stan - of fine sands, silt or mud. The rare at tached gas tro pods, e.g., tial trans port by waves and storms. Thus this as sem blage rep - Diodora graeca, D. italica, Pa tella ferruginea and P. caerula, re sents a shal low fauna thanatocenosis ac cu mu lated close to sug gest the oc cur rence of sporadic gravel and rocky blocks in the coastal line. This is cor rob orated by the oc cur rence in this this shal low-ma rine biotope. The extreme scar city of vermetids de posit of angu lar blocks of deci me ter size, and rounded peb - sug gest the dis per sion of rock blocks and pebbles on which bles. In the Menzel Temime area the ma rine part of the car bon - these gastropoda may build large reefs in ar eas with per ma nent ate unit seems to be de void of Persististombus latus. How ever, ac tive waves and swell as in a few present-day Med i ter ra nean Stramonita haemastoma is fre quent. This spe cies is still alive in lo cal i ties (Fevret and Sanlaville, 1966; Safriel, 1974; Facies and fauna proxies used to reconstruct the MIS 5 and MIS 7 coastal environments in eastern Tunisia 195

Fig. 5A – Persististrombus sp., B–D – Persististrombus latus, E – Semicassis granulata undulata, F – Stramonita haemastoma, G – Conus ventricosus

Dalongeville, 1995, 1996; Vescogni et al., 2008). However, the from Qorba area are also large (Meco, 1977; Gar cia-Talavera, oc cur rence of Persististrombus latus Gmelin (a se nior synonym 1990, 1999; Zazo et al., 1993a, b, 2006; De Turk et al., 1999; of Strombus bubonius Lamarck) and Stramonita haemastoma Avila, 2000; Goy et al., 2003, 2006; Ardovini and Cossignani, (orig i nally at trib uted to the ge nus Thais) in di cates that this Up - 2004; Rolan, 2005; Meco et al., 2011; Montesinos et al., 2014). per Pleistocene fauna in hab ited shal low warm ma rine waters This spe cies re quires warm ma rine waters (with an av er age (Bard et al., 1994; DeTurck et al., 1999; Rilov et al., 2001; Meco, tem per ature of 16°C in Feb ru ary and of 27°C in Sep tem ber). 2002; Torres et al., 2006; 2010; Ramirez et al., 2009; Har ding Meco (1977) noted that cur rent strombids live in quiet marine and Harasewych, 2007; Harzhauser and Kronenberg, 2013; biotopes lo cated near the mouths of rivers. Var i ous modes of Sessa et al., 2013; Muhs et al., 2014). life char ac ter iz ing these di verse molluscs are rec ognized in this In the Q1 sec tion (Fig. 2), bor der ing Chiba Wady, close to part of car bon ate unit: (1) car ni vores, e.g., those be long ing to the bridge and few metres from the road lead ing to Menzel Semicassis, Stramonita and Hexaplex; (2) her biv o rous graz - Temime, the fau nal as sem blage of the lower part of the car bon - ers, e.g., those of Persististrombus, Bittium, Cerithium, Rissoa ate unit is also very diverse with many of the same spe cies ob - and Pa tella (3) sus pen sion feed ers, e.g., those of Turritella, served in the ma rine car bonate unit Q1. This re flects a di verse Donax, Pecten, Ostrea and Chama (Perrier, 1954). The oc cur - and pro duc tive biocenosis, fos sil ised near its orig i nal biotope. rence of sus pen sion feed ers re quires that the waters were con- Ac cu mu la tion was in chan nels where the speed of cur rents stantly renew ed at the bot tom. The her bi vore taxa, as at pres - close to the sea-floor enabled rapid trans por ta tion (Fig. 3C). ent, were likely re lated to a mat of sea weed and/or sea grass of Con se quently, the bi valves com prise ex clu sively sep a rate Zostera and Posidonia the traces of which are not fos sil ised. In valves or debris; en tire shells be ing ab sent. their ab sence of the lat ter, these an i mals are able to swal low In ad di tion, among the gas tro pods, the spe cies large amounts of sand to in gest dis persed cyanobacteria Persististrombus latus is om ni pres ent over the whole Qorba (Goiran, 1990). area, where many entire shells of this ma rine gas tro pod are Anal ysis of this ma rine sands part of the car bon ate unit in pre served. De pending on the age of the ani mal the respec tive the Q1 sec tion shows that bioclasts which are abun dant are shells are more or less large and thick. Their size var ies from 8 mainly of bivalve and gas tro pod shells. As in the Up per Pleis to - to 15 cm in height and 7 to 9 cm in width, without any modi fi ca - cene de pos its of Qorba, the micro fauna is scarce and of low di- tion of spe cific char ac ters. This in di cates that all these shells ver sity. A few speci mens of ben thic foraminifera are rec og - (small or large) be long to one and the same pop ula tion of nized, e.g., Am mo nia beccarii and Elphidium crispum. Persististrombus latus. Com pared to those liv ing to day on the In sum, the sed i men tary fa cies of the car bon ate unit of the shal low-ma rine biotopes of Sen egal, Ga bon, An gola as well as Qorba sec tor is char ac ter is tic of channelized ma rine de pos its around the Cape Verde is lands and the Gulf of Guinea, which (rel atively coarse and well-sorted sands). Thus they cor re - can be reach 100 m in depth, the Up per Pleis to cene spec i mens spond, in all likeli hood, to storm de pos its ac cu mu lated in aban - 196 Amel Chakroun and Dalila Zaghbib-Turki doned chan nels and hol lows within the subtidal zone in which These sands, sug gest that an ex posed bar rier along the beach water was warm and fa vour able to di verse bi valve and gas tro - had formed and ad joined the shal low-ma rine en vi ron ment. pod pop u la tions, es pe cially strombids. In the Hergla area, the quartz-rich unit is char ac ter ized by the same species (Acanthocardia tuberculata, Parvicardium HERGLA SECTIONS exiguum, Ceratoderma edule and Glycymeris nummaria) in the same fre quen cies. Its lithological and fau nal char ac ter is tics The basal part of the car bon ate unit (Fig. 2, Hg1 and Hg2 make it the lat eral equiv alent of the MIS 7 quartz-rich de pos its sec tions) is com posed of sands rich in mol lusc shells with ooids at Menzel Temime and Qorba. How ever, Mauz et al. (2009) de- and rare quartz grains. These sands are inter ca lated with thin fined it as the lower mem ber of the Khnis unit and at trib uted it to co quina beds (Fig. 3Da, b) in which some bi valve spe cies are MIS 5e. Later, Mejri et al. (2012) dated this level at two lo cal i ties dom i nant, e.g., Acanthocardia tuberculata, Ceratoderma edule, in the Sahel (El Hajeb and Hergla) by IRSL and at trib uted it to Parvicardium exiguum, Glycymeris nummaria and Arca noae. MIS 7 in El Hageb whereas no chrono log i cal data have been In this as sem blage the other bi valves are rare (e.g., Arca obtained for the Khnis lo cal ity. tetragona, Chama gryphoides, Loripes lucinalis, Mactra glauca, At Hergla, this quartz-rich de posit is over lain lo cally by len - Mactra stultorum, Callista chione, Spondylus gaederopus, tic u lar clay sug gest ing a la goon with brack ish stag nant-wa ter Tellina planata and Ve nus verrucosa. Gastropoda are also and con ti nen tal de pos its. rare: Semicassis granulata undulata, Columbella rustica, Co- In the Qorba and Hergla ar eas, the lower part of the car bon - nus ventricosus, Hexaplex trunculus, Neverita josephinia, ate unit is rich in molluscs. The as sem blage is very diverse and Persististrombus latus, Turritella communis and Stramonita dom i nated by Acanthocardia tuberculata, Glycymeris haemastoma (Ta ble 2). nummaria, Parvicardium exiguum, Ceratoderma edule and The taphonomy in di cates less tur bu lent shal low-ma rine wa - Arca noae. Nassarius sufflatus and Semicassis granulata ter than in the Cap Bon area. How ever, the or gani za tion of de - undulata are less fre quent. The oc cur rence of ecolog i cal mark - pos its in four se quences with graded bed ding sug gests dis tant ers such as Stramonita haemastoma and Co nus ventricosus storm epi sodes still af fected the Hergla area. (Fig. 5), es pe cially the thermophilic spe cies Persististrombus latus, in di cate shal low warm ma rine water sim i lar to that known ASSEMBLAGE FROM THE MIS 4 AEOLIAN CARBONATE UNIT in the pres ent tropi cal zone. Par tic u larly in the Qorba area, the lower de pos its of the car - In the Qorba as well as in the Hergla area (Fig. 2), the up per bonate unit are or ga nized in three to four se quences fill ing sub - part of the car bonate unit is well-de vel oped. It is com posed of ma rine chan nels. In each se quence, bounded by an erosive eolianites made of car bon ate sands rich in ooids and pel lets. base, sed i men tary clastic and shell el ements (disarticulated, These dunal oolitic sands are rich in var i ous con ti nen tal gas tro - rolled, al tered and even finely broken) are ar ranged in pos i tively pods com posed of He lix spp. and Rumina decollata. How ever graded layers. Each of these se quences sug gests that in the in the Menzel Temime area these coastal dunal de pos its are subtidal aban doned chan nels in ter mit tently ac cu mu lated shells less well-de vel oped or ab sent. and debris dur ing storms as trans gres sion pro ceeded (Chakroun et al., 2009a, b). In Hergla area storms ef fect was mi nor. The up per part of the car bon ate unit, rich in ooids, pel lets, PALAEOENVIRONMENTAL RECONSTITUTION ter res trial gastropoda and root struc tures, cor re sponds to MIS 4 coastal dunal de pos its (Chakroun, 2006; Mauz et al., 2009; The di verse as sem blages from the ar eas stud ied, by their Chakroun et al., 2009a, b). These dunal de pos its are con sis tent struc ture, the eco log i cal sig nif i cance of spe cies con tent and with strong winds act ing on an exposed plat form dur ing the Late their pres er va tion, in form about their hab i tats and their Pleis to cene. They are depos ited dur ing low stand con di tions depositional en vi ron ment (Ta ble 4). (Le Guern, 2005; Frébourg et al., 2008). In the Menzel Temime area, the fauna within the lower part of the quartz-rich unit is very diverse. This biodiversity re flects op ti mal en vi ron men tal con di tions char ac ter iz ing their biotope. PERSISTISTROMBUS LATUS DISTRIBUTION The pres er va tion of many disarticulated bi valve valves dem on - strates that the biocenosis did not un dergo post mor tem trans - In Tuni sia, the ma rine part of the car bonate unit contain ing port. The as sem blage is dom i nated by endobenthic spe cies, Persististrombus latus Gmelin is com monly attrib uted to the iso - liv ing in a loose sub strate com posed of sands or mud. It is also to pic substage 5e or MIS 5.5 (Dubar et al., 2008; Torres et al., char ac ter is tic of a mod er ately ag i tated shal low-ma rine biotope. 2010). This biomarker is fre quent and of ten as so ci ated with Far ther south in the Qorba area, the as sem blage from the Stramonita haemastoma and Nassa circumcincta at many lo- lower part of the quartz-rich unit, at trib uted to MIS 7, is dom i - cal i ties near the north east ern coast line of the Cap Bon Pen in - nated by cardiids and glycymerids, e.g., Acanthocardia sula (Fig. 6B), e.g., near Qorba, Tazaraka and Lebna beach. tuberculata, Parvicardium exiguum, Ceratoderma edule and How ever, in the Menzel Temime area, it is ab sent but Glycymeris nummaria. By contrast Nassarius sufflatus and Stramonita haemastoma and Nassa circumcincta per sist Semicassis granulata undulata are less fre quent. Most of these (Chakroun, 2006). Near the northw est ern coast line of this pen - mol lusc spe cies (Ta ble 4) are of a shal low-ma rine biotope, liv - in sula the Up per Pleis to cene de pos its are dif fer ent. They are ing bur ied in fine sand and mud sub strates. com posed of con sol i dated sands in which molluscs are ab sent Shells are com monly preserved en tire, and debris is rare. but deca pod bur rows (e.g., Callianassa sp.) are com mon. These shells are reg ularly arranged in pla nar laminae. This These lat ter sug gest a supratidal en vi ron ment. sug gests that the biota was af fected by little or no trans port from In the north east lo cal ity of the Cap Bon Pen in sula in the El its biotope. This biotope may be compared to that of a shel tered Haouaria area, the car bonate unit is com posed of sands rich in bay, char ac ter ized by calm-wa ter. Acanthocardia tuberculata and Glycymeris nummaria over lain In the Qorba area, the ma rine part of the quartz-rich unit is by soil, in which pres ent-day ploughing exhumes rare spec i - over lain by fine sands with abun dant root traces (Fig. 3Db). mens of Persististrombus latus (Fournet, 1981; Chakroun, Facies and fauna proxies used to reconstruct the MIS 5 and MIS 7 coastal environments in eastern Tunisia 197

Ta ble 4

Tax o nomic com po si tion and ecol ogy of spec i mens iden ti fied in the Up per Pleis to cene de pos its

TAXA ECOLOGICAL CHARACTERISTICS MT QRB HRG KN(a) BIVALVIA Sub strate/tier ing/mo bil ity/feed ing Sand, mud or gravel/epifaunal/facultatively mobile/sus - Acanthocardia tuberculata (Linnaeus, 1758) pen sion feeder Barbatia barbata Linnaeus, 1758 Hard/epifaunal/sta tion ary, at tached/sus pen sion feeder Arca noae Linnaeus, 1758 Hard/epifaunal/sta tion ary, at tached/sus pen sion feeder Arca tetragona PolI, 1795 Hard/epifaunal/sta tion ary, at tached/sus pen sion feeder Sand, mud and gravel/endobenthic/mo bile/sus pen sion Ceratoderma edule (Linnaeus, 1758) feeder Callista chione (Linnaeus, 1758) Sand/infaunal/facultatively mo bile/sus pen sion feeder Chama gryphoides Linnaeus, 1758 Hard/epifauna/sta tion ary, at tached/sus pen sion feeder Sand and mud/infaunal/facultatively mo bile/sus pen sion Chamelea gallina (Linnaeus, 1758) feeder Sand/infaunal/facultatively mobile, creeping/sus pen s ion Dosinia exoleta Linnaeus, 1758 feeder Sands and mud/infaunal/facultatively mobile/sus pen sion Donax trunculus Linnaeus, 1758 feeder Sand/infaunal/facultatively mo bile, creep ing/sus pen sion Dosinia lupinus (Linnaeus, 1758) feeder Sand and mud/ infaunal/ facultatively mobile/ sus pension Eastonia rugosa (Helbing, 1779) feeder Ensis ensis Linnaeus, 1758 Sand or mud/infaunal/stationnay/sus pen sion feeder Gastrana fragilis (Linnaeus, 1758) Sand/ infaunal/facultatively mo bile/de posit feeder Glycymeris nummaria (Linnaeus, 1758) Sand or gravel/semi-infaunal/creep ing/sus pen sion feeder Sand and gravel/deep infaunal/facultatively mo- Loripes lucinalis (Lamarck, 1818) bile/chemosymbiotic Mactra stultorum (Linnaeus, 1758) Sand/infaunal/facultatively mobile/sus pen sion feeder Mactra glauca Born, 1778 Sand/infaunal/facultatively mobile/sus pen sion feeder Pitar rudis (Poli, 1795) Sand/infaunal/facultatively mobile/sus pen sion feeder Modiolus barbatus (Linnaeus, 1758) Hard/semi-infaunal/station ary/sus pen sion feeder Ostrea edulis Linnaeus, 1758 Hard/epifaunal/immo bile attached/sus pen si on feeder Parvicardium exiguum Gmelin, 1791 Sand/epifaunal/facultatively mo bile/sus pen sion feeder Pecten jacobaeus (Linnaeus, 1758) Sand/epifaunal/facultatively mo bile/sus pen sion feeder Polititapes rhomboides (Pennant, 1777) Mixed/infaunal/facultatively mo bile/sus pen sion feeder Pseudochama gryphina (Lamarck, 1819) Hard//epifaunal/im mo bile at tached/sus pen sion feeder Solecurtus strigilatus (Linnaeus, 1758) Sand/infaunal/facultatively mo bile/sus pen sion feeder Spondylus gaederopus Linnaeus, 1758 Hard/epifaunal/im mo bile at tached/sus pen sion feeder Sand, muddy gravel and clay/infaunal/facultatively mo- Ruditapes decussatus (Linnaeus, 1758) bile/sus pen sion feeder Sand, muddy gravel and clay/infaunal/facultatively mo- Tellina planata Linnaeus, 1758 bile/suspen sion feeder Sands, muddy gravel or clay/ infaunal/ facultatively mo- Venus verrucosa Linnaeus, 1758 bile/suspen sion feeder GASTROPODA Sea grass sys tem/epibenthic/mo bile/grazer, de posit Bittium reticulatum (Da Costa, 1778) feeder Cerithiopsis min ima (Brusina, 1865) Hard/epifaunal/mo bile/ectoparasite and car ni vore Live and gaze on sponge/epifaunal/mo bile/de posit Cerithiopsis tubercularis (Montagu, 1803) feeder, car ni vore Cerithium vulgatum Bruguiere, 1792 Mixed/epifaunal/mo bile/her bi vore Columbella rustica (Linnaeus, 1758) Hard/ epifaunal/creep ing/om ni vore-grazer, Her bi vore Co nus ventricosus Gmelin, 1791 Mixed/epifaunal/creep ing/scav en ger, pred a tor Cyclope neritea (Linnaeus, 1758) Sands and mud/epifauna/slow-mov ing/car ni vore Hard/epifaunal/attached, facultatively mo- Diodora italica (Defrance, 1820) bile/ectoparasites and car ni vore, her bi vore Hard/epifaunal/attached, facultatively mo- Diodora graeca (Linnaeus, 1758) bile/ectoparasites and car ni vore, her bi vore Hexaplex trunculus (Linnaeus, 1758) Mixed/epifaunal/creep ing/car ni vore, pred a tor,scav en ger Monophorus perversus (Linnaeus, 1758) Mixed/epifaunal/mo bile/ectoparasites and car ni vore Sandy and grav els/epifaunal/ slow-mov ing low-level/car - Nassarius reticulatus (Linnaeus, 1758) ni vore, scav en ger Sandy and grav els/epifaunal/slow-mov ing low-level/car ni - Nassarius sufflatus (Gould, 1860) vore, scav en ger 198 Amel Chakroun and Dalila Zaghbib-Turki

Tab. 4 cont.

ECOLOGICAL CHARACTERISTICS TAXA MT QRB HRG KN(a) Substrate/tier ing/mo bil i ty/feeding Neverita josephinia Risso, 1826 Sand/infaunal/slow-mov ing/car ni vore, pred a tor Naticarius stercusmuscarum (Gmelin, 1791) Sand or mud/infaunal/slow-mov ing/car ni vore, scav en ger Pa tella caerula Linnaeus, 1758 Hard/epifaunal/facultatively mo bile, at tached/her bi vore Pa tella ferruginea Gmelin, 1791 Hard/epifaunal/facultatively mo bile, at tached/her bi vore Pa tella vulgata Linnaeus, 1758 Hard/epifaunal/facultatively mo bile, at tached/her bi vore Persististrombus latus (Gmelin, 1791) Sand/epifaunal/creep ing/her bi vore, om ni vore-grazer Rissoa membranacea (Adams, 1800) Mixed/epifaunal/mo bile/her bi vore, grazer Semicassis granulata undulata (Gmelin, 1791) Mixed/epifaunal/creep ing, grazer/predatoroy, car ni vore Stramonita haemastoma (Linnaeus, 1758) Hard/epifaunal/creep ing, pred a tor/scav en ger, car ni vore Tarantinaea lignaria (Linnaeus, 1758) Hard/epifaunal/herbiers/car niv o rous. Thylacodes arenarius (Linnaeus, 1758) Hard/sta tion ary, at tached/infaunal/sus pen sion feeder Turritella communis Risso, 1826 Sand/epifaunal/facultatively mo bile/sus pen sion feeder He lix spp. Mixed (ter res trial), epifaunal/free/her biv o rous Rumina decollata Linnaeus, 1758 Mixed/ter res trial snail/car ni vore/pred a tor. ECHINOIDEA Echinocardium cordatum (Pen nant, 1777) Sand/infaunal/slow-mov ing/de posit feeder-detritivore FORAMINIFERA Am mo nia beccarii (Linnaeus, 1758) ben thic sta tion ary semi-infaunal/her bi vore Am mo nia tepida (Cushman, 1926) bentic/sta tion ary semi-infaunal/her bi vore Asterigerinata mamilla (Wil liam son, 1858) ben thic/sta tion ary semi-infaunal/om ni vore Elphidium crispum (Linnaeus, 1758) ben thic/sta tion ary semi-infaunal/om ni vore Nonion sp. ben thic/sta tion ary semi-infaunal/om ni vore Textularia sp ben thic/sta tion ary infaunal/detritivore MAMMELS Phacochoerus aethiopicus (Pallas, 1766) ground dwell ing om ni vore Gazella atlantica Bourguinat, 1870 ground dwelling grazer-browser Alcelaphus buselaphus (Pallas, 1766) ground dwelling grazer Equus mauritanicus Pomel, 1897 ground dwelling grazer Ceratotherium simum (Burchell, 1817) ground dwelling grazer Hystrix cristata Linnaeus, 1758 semi fossorial her bi vore-frugivore

MT – Menzel Temime, QRB – Qorba, HRG – Hergla, KN – Khnis, (a) – addi tional data from (Mahmoudi, 1986); grey – spe cies oc cur rence; for the fau nal de ter mi na tion and the eco log i cal char ac ter is tics, we re ferred to Abbott (1960), Perrier (1954), Fekih (1975), Meco, (1977) and Sessa et al. (2013); taxo nomic at tri butions were updated by using the da tabase: the World Reg is ter of Marine Spe cies

2006). Far ther tow ards the south, in the Kelibia area, this Mogods and Cap Blanc coast line Persististrombus latus spe - biomarker is usu ally asso ci ated with or replaced by Stramonita cies has been never re ported (Solignac, 1927; Miossec, 1977; haemastoma. Chakroun, 2006). However, east of Bizerte city near the R’Mel Along the rest of the east ern coast line of Tuni sia (Fig. 6C), beach it is spo rad i cally present in car bon ate bioclastic sands Persististrombus latus is of ten pres ent in the car bon ate unit, es - (Ben Ayed et al., 1978). In con trast, it is more frequent in the pe cially near the beaches of Hergla, Monastir, Khnis, Mahdia, Raraf area (Chakroun et al., 2016). Rejiche, Chebba, Kerkennah and Djerba (Castany, 1954, Else where in the Up per Pleis to cene de pos its sit u ated 1955, 1956; Paskoff and Sanlaville, 1976; Oueslati, 1980; around the Med i ter ra nean Sea, the Persististrombus latus Kamoun, 1981; Oueslati et al., 1982; Mahmoudi, 1986; Jedoui, show a dif fer ent dis tri bu tion. In the Lev ant, it was re ported in 2000; Chakroun, 2006; Chakroun et al., 2009a). It is repre - Leb a non (Fleisch and Gigout, 1966; Fleisch and Sanlaville, sented by large spec i mens with large tuber cles, also found by 1967; Sanlaville, 1969, 1978; Fleisch et al., 1971), in North Mahmoudi (1986). Such forms are rem i nis cent of their coun ter - Haifa (Issar and Kafri, 1972; Sivan et al., 1999), in Cyprus es pe - parts liv ing to day in the Gabon coastal area to a depth of 4 to 8 cially in Larnaca and west to Coral Bay (Poole et al., 1990). In m (Ber nard, 1984). Far ther tow ards the south, near the coast - Greece, the Sen ega lese fauna com posed of Loripes lucinalis, line of Gabes and Djerba Is land, the car bon ate unit is rich in Co nus testudinarius, Cantharrus viverratus and Stramonita strombids, ooids, pel lets of crus ta ceans and bioclasts haemastoma and in clud ing Persististrombus latus was sig naled (Castany, 1955; Paskoff and Sanlaville, 1977). This fa cies in di - in many local i ties in bor der of the south coast of Crete cates a warm and agi tated plat form (Lucas, 1955; Purser, (Keraudren et al., 2000) and in It aly (Hearty et al., 1986a, b; 1980; Chakroun, 2006). Balescu et al., 1997). It is pres ent in the isoto pic stage 5 ter - In north ern Tu ni sia (Fig. 6A), only Stramonita haemastoma races along the south ern Calabria (Dumas et al., 1988, 2002), with out strombids was re ported in Pleis to cene de pos its of the in bor der of the Ionian coast and in Crotone Pen in sula (Nalin et Galite Is land (Issel, 1880). Tow ards the south along the al., 2012) as well as Sic ily (Gignoux, 1913; Antonioli et al., Facies and fauna proxies used to reconstruct the MIS 5 and MIS 7 coastal environments in eastern Tunisia 199

Fig. 6. Distribution map of Persististrombus latus on the Tunisian coast 200 Amel Chakroun and Dalila Zaghbib-Turki

Fig. 7. Distribution map of the Persististrombus latus sites on the Mediterranean coast Sources of data: Syria (Sanlaville, 1979); North Haïfa (Issar and Kafri, 1972; Sivan et al., 1999); Leb a non (Fleisch and Gigout, 1966); Cy- prus (Poole et al., 1990); Greece (Keraudren et al., 2000); Italy (Gignoux, 1913; Ozer et al., 1980; Hearty et al., 1986a, b; Dumas et al., 2002; Antonioli et al., 2006; Nalin et al., 2012); Tuni sia : pres ent work; France (Valensi et al., 2007; Dubar et al., 2008); Spain (Hillaire-Mar cel et al., 1996; Zazo, 1999; Zazo et al., 2003; Goy et al., 2006); Al ge ria (De Lamothe, 1911); Morocco (Guillemin, 1976; Hearty, 1986; Sbai, 2008; Muhs, 2015)

2006). In Sar dinia strombids are rare (Ozer et al., 1980; Kin dler Con se quently, this dis tri bu tion of the Sen e ga lese fauna es - et al., 1997; Le Guern, 2005). In this is land the Up per Pleis to - pe cially Persististrombus latus (Fig. 7) in the circum-Med i ter ra - cene as sem blage is mainly com posed of Pa tella ferruginea and nean ar eas (in North Af rica and Eu rope) al lows the re con struc - Co nus testudinarius (Lecca and Carboni, 2007). In Spain tion of the shal low warm ma rine zone dis tri bu tion dur ing the Persististrombus latus was found in Almeria, Alicante and Late Pleis to cene. Persististrombus latus is a shal low-ma rine Mallorca (Hillaire-Mar cel et al., 1986; Zazo and Goy, 1989; thermophilic spe cies liv ing nearby sheltered beaches. Hillaire-Mar cel et al., 1996; Zazo, 1999). Its first ap pear ance in Only while these ecolog i cal con di tions are met this Almeria is as so ci ated to the MIS 7a (Zazo, 1999). biomarker thrived. The spe cies Persististrombus latus or its se- In southeast ern of France, be tween Nice and Menton, the nior synonym Strombus bubonius were fre quently sig naled in dis tri bu tion of the Sen e ga lese fauna in clud ing Persististrombus many Med i ter ra nean coastal Tyrrhenian de pos its (Gignoux, latus was used to recon struct the palaeocoast line char ac ter iz - 1911). ing the Tyrrhenian period (Bonifay, 1961; Valensi et al., 2007; In the circum Med i ter ra nean Sea the Sen e ga lese Fau nal Dubar et al., 2008). The palaeoshore to wards the south of Nice Com plex as so ci a tion in clud ing Persististrombus latus and other in cluded Grimaldi caves in Ital ian Liguria (Cave of the Prince, mol lusc spe cies in di cates sea wa ter warm ing dur ing inter gla cial Barma Grande), Mo naco and Cap Ferrat (Bonifay, 1975). In ep i sodes along the Pleis to cene, which cor re sponded to pe ri ods north ern Mo rocco, the Persististrombus latus oc curs in the of high sea level. In fact, cur rent P. latus is re stricted to the East - Pleis to cene ma rine de pos its at El Hoceima (Guillemin, 1976; ern At lan tic Re gion, oc cur ring in an area from the Rio d'Oro to Hearty et al., 1986b; Sbai, 2008; Muhs, 2015). In Al geria the An gola and lim ited in the north by the Canary Current and in the most im por tant fos sil de pos its of Persististrombus latus are lo - south by the Benguela Cur rent. Its north ern dis tri bu tion bor ders cated in Gulf of Oran, Gulf of Arzew and Gulf of Mostaganem are the Cape Verde Is lands (Kreipl et al., 1999). An av er age (De Lamothe, 1911). tem per ature of 25°C fa vours this gas tro pod; how ever, it tol er - Facies and fauna proxies used to reconstruct the MIS 5 and MIS 7 coastal environments in eastern Tunisia 201 ates a tem per ature range from 16 to 31°C. Its av er age life span lucinalis, pres ently known to be able to live in con fined biotopes of 3–5 years is char ac ter ized by a cy clic growth rate, de pend ing in which dis solved ox ygen is low and sa lin ity is vari able, may in- on the sea son al ity of the sea sur face tem per ature. P. latus is a di cate la goon with stagnant and con fined water. gre gar i ous spe cies can toned to shal low-wa ters (Meco, 1977; An up per ae olian quartz-rich term is as so ci ated to MIS 7 Mendes-Lopes, 2002). It is known in sandy-muddy bot toms and and MIS 5e la goon de pos its of unit de posi tion these ma rine or sea grass meadow s and fol lows detritivorous and veg e tar ian la goonal en vi ron ments be came ex posed and coastal dunes feed ing strat e gies (Lozac'hmeur and Mascarenhas, 1985), typi - were de vel oped. It had been fixed by more or less dense veg e - cally in waters with normal sa lin ity (Guerreiro, 1994; Guerreiro ta tion and con tained con ti nen tal snails (He lix spp. and Rumina and Reiner, 2000; Mendes-Lopes, 2002). However, in the Gulf de col lata). The abun dance of quartz grains in this ae olian part of Guinea, Meco (1977) found P. latus rep re sen ta tives in zones sug gests rel a tive hu mid con di tions along the palaeoshore. with fresh wa ter in put and con se quently lower sa lin ity. Al though The depo si tion of the lower part of the car bonate unit, rich in P. latus in habits soft bot toms, and may be sheltered by rocky ma rine fauna of MIS 5, in the Cap Bon area as well as the blocks. Hergla area in di cates marine trans gres sion. In this shal low-sea Com par ing Pleis to cene and Pres ent dis tri bu tion of P. latus there thrived di verse shal low-ma rine spe cies, e.g., bi valves: we note that dur ing the Late Pleis to cene this warm species was Acanthocardia tuberculata, Ceratoderma edule, Parvicardium can toned to circum Medi ter ra nean Sea. At pres ent this spe cies exiguum, Glycymeris nummaria, Arca noae, A. tetragona, as well as its con generic spe cies, has become re stricted to trop i - Chama gryphoides, Loripes lucinalis, Mactra coralina, M. cal seas. It is clear that this thermophilic spe cies mi grated dur ing glauca, Callista chione, Spondylus gaederopus, Tellina the gla cial pe riod suc ceed ing the 5e substage in or der to find planata, Ve nus verrucosa. Gas tro pods are less fre quent with again warm water (warmer than in cur rent Med i ter ra nean Sea). Semicassis granulata undulata, Columbella rustica, Co nus ventricosus, Hexaplex trunculus, Nassarius sufflatus, Neverita josephinia, Persististrombus latus, Turritella communis, SUMMARY AND CONCLUSIONS Stramonita haemastoma. Within this as sem blage thermophilic spe cies are fre quent as in the Sen ega lese fauna. These Pleis to cene (MIS 7, MIS 5e and MIS 4) de pos its ex - Persististrombus latus is es pecially common. Its pres ence is in - posed along the east ern coast of Tuni sia, by their di verse facies dic ative of cli mate warm ing. The group ing of de pos its and and fauna con tents, pro vide valu able in for ma tion about en vi - fauna into four graded se quences sug gests that the warmer cli - ron ments prevail ing dur ing this pe riod. Tak ing into ac count the mate in cluded storm ep i sodes. palaeo eco logi cal re quire ments of dif fer ent spe cies com pos ing As the up per part of the car bonate unit was depos ited ma - the suc ces sive fau nal as sem blages, spe cies rich ness, spe cies rine re gres sion exposed the en tire coast line in clud ing the pre vi - rel a tive abun dance and pres er va tion sta tus of the dif fer ent ous subtidal en vi ron ment. In the backshore area large dunes spec i mens, the Pleis to cene palaeoenvironments are re stored. were de vel oped, at trib uted to MIS 4, sug gest ing strong wind ac- These as sem blages are rep re sen ta tive of bi o log i cal com mu ni - tion. ties which have been af fected by more or less dis tant trans por - In sum, the Mid-Up per Pleis to cene fa cies and fauna sug - ta tion. In all the cases these marine com mu ni ties were de vel - gest dif fer ent en vi ron ments related in space and over time: (1) oped dur ing an inter gla cial warm pe riod. Any change in the as - a subtidal envi ron ment with shel tered bay; (2) a la goonal en vi - sem blages tra duces change in com mu ni ties. These changes ron ment with oligospecific fauna, (3) a subtidal en vi ron ment re - would be re lated to cli mate and en vi ron men tal fluc tu a tions. cord ing storm ac tion in sub ma rine chan nels, (4) a shore line Thus, fa cies and fau nal as sem blage char ac ter is tics in the with coastal dunes re sult ing from strong wind action. basal term of the quartz-rich unit (Douira For ma tion), at trib uted to the MIS 7, in di cates quite ma rine shal low envi ron ment in Cap Ac knowl edge ments. We are grate ful to E. Davaud for the Bon Pen in sula. Well-pre served shells of abun dant cardiids, week of field re search in Hergla. This manu script was im proved glycymerids, arcids, lucinids and donacids sug gest shal low and by the detailed and thought ful com ments of the Ed i tor quite ma rine water char ac ter is tic of a shel tered bay or part of la - W. Granoszewski, the review er P. Sümegi, and an anon ymous goon con nected with the sea. re viewer. We would like to ac knowl edge the writer M. Dick ens The quartz-rich de pos its (Khnis For ma tion) iden ti fied in who kindly review ed the Eng lish lan guage of this manu script. Hergla and at trib uted to the MIS 5e show the same spe cies as Finaly, we would like to thank Dr. J. Jaballah for his support in the Cap Bon (MIS 7) de pos its. In deed, the pres ence of Loripes infography.

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